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Rearing trials of Halla parthenopeia under laboratory conditions (Polychaeta: Oenonidae) Inas H. Osman , Howaida R. Gabr, Salah Gh. El-Etreby Marine Science Department, Faculty of Science, Suez Canal University, Ismailia, Egypt abstract article info Article history: Received 14 February 2009 Received in revised form 29 October 2009 Accepted 29 October 2009 Keywords: Biochemical composition Growth Halla parthenopeia Preferred food species Suez Canal A small scale attempt to maintain and rear the worm Halla parthenopeia in laboratory conditions was conducted. Five bivalve species (Paphia undulata, Cerastoderma glaucum, Venerupis pullastra, Ruditapes decussata, and Gafrarium pectinatum) were used to investigate preferred food item, feeding rate and growth of the worm. Halla parthenopeia are capable of using different types of clams, although they grew better with Paphia undulata and C. glaucum as food items. The highest average daily predation rates in case of P. undulata as a prey were 1.73, 2.13 and 2.57 individuals per predator per day for small, medium and large groups of H. parthenopeia, respectively. The daily predation rate on C. glaucum was low with an average: 0.50, 0.63 and 0.73 individuals per predator per day for the small, medium and large worm groups, respectively. The daily growth rate of H. parthenopiea increased when it was fed P. undulata (average: 0.083, 0.071 and 0.038 g/day for small, medium and large worm groups respectively), compared to an average of 0.044, 0.034 and 0.020 g/ day for small, medium and large worm groups, respectively, when worms were fed with C. glaucum. The biochemical composition of three different sizes of the worm was also determined. Protein was the highest biochemical constituent with an average of 51% of the dry weight, followed by lipids with an average of 25.88% of the dry weight; meanwhile carbohydrate was present at an average of 20.72%. Our ndings indicate that growth of H. parthenopeia can be improved when fed with a suitable prey item and suggest that it is feasible to successfully culture this protein-rich worm in captivity. © 2009 Elsevier B.V. All rights reserved. 1. Introduction The worm Halla parthenopeia (Delle Chiaje, 1828) belongs to the family Oenonidae and it is widely distributed throughout the Suez Canal. It is a large polychaete which can reach up to 90 cm in length (Osman, 2007). Halla parthenopeia is exploited for shing bait, especially for sea bream and as a result, a thriving collecting business for bait was developed along Suez Canal. So, this worm could be a promising candidate for commercial aquaculture. It is distributed from the eastern Mediterranean (Selim, 1978; Abd El-Naby, 2005) through the Suez Canal (Barbary, 1992) across to The Red Sea (Belal, 2001). It lives in ne to coarse sediment in the sublittoral water of the Suez Canal (Osman, 2007) and feeding mainly on bivalve molluscs. In Egypt, little is known about this species, particularly with regards to reproduction and culture (Rouse and Pleijel, 2001; Osman et al., in press). The potential commercial value of Polychaeta is not limited to their use as shing baits; another value is being discovered with growth of the aquaculture industries for the production of nsh and crustacea (Costa and Bruxelas, 1989). Commentators on the aquaculture industry on a world scale identify the current shortfall in the production of juveniles as a limiting factor, as a potential danger to expansion of the aquaculture industry, and a major cause of non- sustainability. It has been suggested that the supply of polychaetes as food for brood stocks could help to alleviate this problem (Olive, 1999). This commercial value arises from the high palatability of many species to commercially farmed sh. Furthermore, their lipid contents may provide a source of essential polyunsaturated fatty acids (PUFAs) (Luis and Passos, 1995) that are required for successful production of high quality juveniles of both nsh and crustaceans (Olive, 1999). Polychaeta could also be used for the improvement of diets in hatcheries and larval rearing units, such as Penaeus monodon shrimp. The culture of polychaetes has recently become a very important area due to their increased market price and their potential role that may have for the improvement of brood stock output in many sectors of the world aquaculture industry (Olive, 1999). In the present study, a small scale attempt to maintain and rear H. parthenopeia in laboratory was conducted. The preferable food items, feeding rate and growth were compared using different species of bivalves. This will help to determine the most suitable bivalve species feasible in the culture of this polychaete on a commercial scale. The focus of the study was on foraging behavior and suitable conditions of water temperature, salinity, pH and suitable type of substrate. The biochemical composition (Protein, Lipids and Carbohy- drates) of H. parthenopeia was also assessed to evaluate its nutritive Journal of Experimental Marine Biology and Ecology 383 (2010) 17 Corresponding author. Tel.: +20 2 0103618559. E-mail address: [email protected] (I.H. Osman). 0022-0981/$ see front matter © 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.jembe.2009.10.016 Contents lists available at ScienceDirect Journal of Experimental Marine Biology and Ecology journal homepage: www.elsevier.com/locate/jembe

Rearing trials of Halla parthenopeia under laboratory conditions (Polychaeta: Oenonidae)

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Page 1: Rearing trials of Halla parthenopeia under laboratory conditions (Polychaeta: Oenonidae)

Journal of Experimental Marine Biology and Ecology 383 (2010) 1–7

Contents lists available at ScienceDirect

Journal of Experimental Marine Biology and Ecology

j ourna l homepage: www.e lsev ie r.com/ locate / jembe

Rearing trials of Halla parthenopeia under laboratory conditions(Polychaeta: Oenonidae)

Inas H. Osman ⁎, Howaida R. Gabr, Salah Gh. El-EtrebyMarine Science Department, Faculty of Science, Suez Canal University, Ismailia, Egypt

⁎ Corresponding author. Tel.: +20 2 0103618559.E-mail address: [email protected] (I.H. Osman).

0022-0981/$ – see front matter © 2009 Elsevier B.V. Aldoi:10.1016/j.jembe.2009.10.016

a b s t r a c t

a r t i c l e i n f o

Article history:Received 14 February 2009Received in revised form 29 October 2009Accepted 29 October 2009

Keywords:Biochemical compositionGrowthHalla parthenopeiaPreferred food speciesSuez Canal

A small scale attempt to maintain and rear the worm Halla parthenopeia in laboratory conditions wasconducted. Five bivalve species (Paphia undulata, Cerastoderma glaucum, Venerupis pullastra, Ruditapesdecussata, and Gafrarium pectinatum) were used to investigate preferred food item, feeding rate and growthof the worm. Halla parthenopeia are capable of using different types of clams, although they grew better withPaphia undulata and C. glaucum as food items. The highest average daily predation rates in case of P. undulataas a prey were 1.73, 2.13 and 2.57 individuals per predator per day for small, medium and large groups of H.parthenopeia, respectively. The daily predation rate on C. glaucum was low with an average: 0.50, 0.63 and0.73 individuals per predator per day for the small, medium and large worm groups, respectively. The dailygrowth rate of H. parthenopiea increased when it was fed P. undulata (average: 0.083, 0.071 and 0.038 g/dayfor small, medium and large worm groups respectively), compared to an average of 0.044, 0.034 and 0.020 g/day for small, medium and large worm groups, respectively, when worms were fed with C. glaucum. Thebiochemical composition of three different sizes of the worm was also determined. Protein was the highestbiochemical constituent with an average of 51% of the dry weight, followed by lipids with an average of25.88% of the dry weight; meanwhile carbohydrate was present at an average of 20.72%. Our findingsindicate that growth of H. parthenopeia can be improved when fed with a suitable prey item and suggest thatit is feasible to successfully culture this protein-rich worm in captivity.

l rights reserved.

© 2009 Elsevier B.V. All rights reserved.

1. Introduction

The worm Halla parthenopeia (Delle Chiaje, 1828) belongs to thefamily Oenonidae and it is widely distributed throughout the SuezCanal. It is a large polychaete which can reach up to 90 cm in length(Osman, 2007). Halla parthenopeia is exploited for fishing bait,especially for sea bream and as a result, a thriving collecting businessfor bait was developed along Suez Canal. So, this worm could be apromising candidate for commercial aquaculture. It is distributedfrom the eastern Mediterranean (Selim, 1978; Abd El-Naby, 2005)through the Suez Canal (Barbary, 1992) across to The Red Sea (Belal,2001). It lives in fine to coarse sediment in the sublittoral water of theSuez Canal (Osman, 2007) and feeding mainly on bivalve molluscs. InEgypt, little is known about this species, particularly with regards toreproduction and culture (Rouse and Pleijel, 2001; Osman et al., inpress).

The potential commercial value of Polychaeta is not limited to theiruse as fishing baits; another value is being discovered with growth ofthe aquaculture industries for the production of finfish and crustacea(Costa and Bruxelas, 1989). Commentators on the aquacultureindustry on a world scale identify the current shortfall in the

production of juveniles as a limiting factor, as a potential danger toexpansion of the aquaculture industry, and a major cause of non-sustainability. It has been suggested that the supply of polychaetes asfood for brood stocks could help to alleviate this problem (Olive,1999). This commercial value arises from the high palatability ofmany species to commercially farmed fish. Furthermore, their lipidcontentsmay provide a source of essential polyunsaturated fatty acids(PUFAs) (Luis and Passos, 1995) that are required for successfulproduction of high quality juveniles of both finfish and crustaceans(Olive, 1999). Polychaeta could also be used for the improvement ofdiets in hatcheries and larval rearing units, such as Penaeus monodonshrimp. The culture of polychaetes has recently become a veryimportant area due to their increasedmarket price and their potentialrole that may have for the improvement of brood stock output inmany sectors of the world aquaculture industry (Olive, 1999).

In the present study, a small scale attempt to maintain and rearH. parthenopeia in laboratory was conducted. The preferable fooditems, feeding rate and growth were compared using different speciesof bivalves. This will help to determine the most suitable bivalvespecies feasible in the culture of this polychaete on a commercialscale. The focus of the study was on foraging behavior and suitableconditions of water temperature, salinity, pH and suitable type ofsubstrate. The biochemical composition (Protein, Lipids and Carbohy-drates) of H. parthenopeia was also assessed to evaluate its nutritive

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value and to determine its commercial potential in aquaculture. Thisresearch presents the first step towards farming H. parthenopeiasuccessfully.

2. Materials and methods

2.1. Collection of samples and preparation of the aquaria

Specimens of H. parthenopeia were collected in June 2006 fromLake Timsah at Ismailia city along Suez Canal area (29°56′13.84″N and32°34′4.85″E and 31°16′4.49″N and 32°19′12.09″E)(Fig. 1). Wormswere collected by hand while snorkeling at depths ranging between 1

Fig. 1. A map showing Suez Can

and 5 m, with water clarity ranging from 40 cm to 60 cm. They weretransferred to an experimental tank in the laboratory and starved for aweek before the start of the experiment to exclude the influence ofpast feeding experience (Saito et al., 2000).

The most abundant species of bivalves (Paphia undulata (3–3.5 cm), Cerastoderma glaucum (2–2.6 cm), Venerupis pullastra (1.4–3.5 cm), Ruditapes decussata (1.4–1.8 cm), and Gafrarium pectinatum(1.8–2.2 cm)) were collected from Lake Timsah to be used as preys inthe feeding experiments and were stored in three aerated tanks of90 L each. Each species was sorted depending on its available sizerange and was classified to small, medium and large sized. Each tankcontained five different species of the same size group.

al and related water bodies.

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3I.H. Osman et al. / Journal of Experimental Marine Biology and Ecology 383 (2010) 1–7

The experimental tanks were 13 L each of rectangular polyethyl-ene plastic with dimensions 35 cm×25 cm×15 cm containing a layerof 7 cm sediment (30% gravel, 20% sand, 50% mud). The salinity wasmaintained at 35 ppt±1 ppt. The temperature was thermostaticallycontrolled and remained constant at 25 °C by placing immersibleaquarium heaters in each tank, while pH was kept close to 7.8. Theexperimental tanks were connected to a re-circulating system,comprising a mechanical filter, biological filter and a 30 L sump.

All tanks were covered to minimize evaporation of the seawater tomaintain the salinity and to prevent the effect of strong light on theworms. The cover provided complete darkness for the worms exceptfor the periods when prey shells were changed.

2.2. Prey selection and predation rate

Nine individuals of H. parthenopeia were divided into three sizegroups, small (average length 23.9 cm, 7.5–10.5 g), medium (averagelength 34.6 cm, 15.5–18.5 g) and large (average length 43.8 cm, 24.5–29.5 g). Each group had three individuals, which were placed in aseparate aquarium. The experiment lasted for 30 days then repeatedfor three times with different worms of the same size groups andbivalves.

A mixture of five individuals of each of the five prey species wasintroduced to each experimental tank (multiple-choice trials). Thepreys were dispersed over the sediment. It was taken into consider-ation that each prey size group was used to feed the correspondingsize of worms that is to exclude the effect of prey size during preyselection. Tanks were inspected on daily basis and the consumed preywas replaced with a new one of the same species and size. Over aperiod of 30 days, the number of prey consumed by each size groupwas counted and the preferred species was determined. Results ofthree replicates were averaged and the number of prey eaten perindividual worm per day was calculated.

To confirm the results of the preferred bivalve species, anotherexperiment with two alternative trials (two prey species only wereoffered in each tank) was carried out. . Each trial lasted for a period of10 days and the number of prey eaten by each size groupwas counted.Results of ten replicates were averaged.

Table 1Preference and predation rate of three sized groups of Halla parthenopeia for differentprey species.

Prey Predatorsize

Predation rate ofH. parthenopeiaper 30 days

Dailypredationrate

Species Shell length(cm)

Number SD ± %

Paphia undulata b3.0 Small 156 5.4 70.3 1.733.0–3.5 Medium 192 6.9 71.1 2.13N3.5 Large 231 10.2 70.0 2.57

Cerastoderma glaucum b2.2 Small 45 2.5 20.3 0.502.2–2.6 Medium 57 3.1 21.1 0.63

2.3. Growth rate

As a result of the above trials, the most two favorite prey toH. parthenopeiawere selected to test their impact on the growth of theworm. Eighteen individuals of H. parthenopeia were divided betweensix experimental rearing tanks. This allowed two treatment groups tobe tested in triplicate with three animals per tank. The first group wasfed on Paphia undulatawhile the second group was fed on C. glaucum.An equal number of each prey species was offered to each tank. All thetanks were inspected in a daily basis and the remains of the ingestedpreys were removed and replaced by live ones. The experiment lastedfor 60 days.

The total wet weight of each worm was recorded at the beginningand the end of the trial. The daily growth rate (mg/day) wascalculated for each worm as the difference between the final andthe initial body weight divided by the number of days. The results arepresented as averages for each size group.

N2.6 Large 66 6.2 20.0 0.73Venerupis pullastra b1.4 Small 12 1.1 5.4 0.13

1.4–1.8 Medium 9 0.5 3.3 0.10N1.8 Large 18 2.1 5.5 0.20

Ruditapes decussata b1.4 Small 9 0.8 4.1 0.101.4–1.8 Medium 12 0.2 4.4 0.13N1.8 Large 15 1.5 4.5 0.17

Gafrarium pectinatum b1.8 Small 0 0 0.0 0.001.8–2.2 Medium 0 0 0.0 0.00N2.2 Large 0 0 0.0 0.00

2.4. Feeding behavior

Throughout the experiments, the worms were observed daily forfeeding behavior as well as general vitality. At random times duringthe day, focal sampling was used to observe an individual for a periodof 1 h. Continuous recordings of that individual behavior and itsinteraction with the prey was made.

2.5. Biochemical composition analysis

Eighteen specimens of H. parthenopeiawere used to determine thebiochemical composition. The biochemical composition was deter-mined in relation to weight regardless of sex due to the difficulty ofsexual dimorphism. Samples were weighed and separated into threegroups; small (5–12 g), medium (12–16 g) and large (16–32 g), thenplaced in sealed plastic bags and stored in a deep freezer for furtheranalysis. Moisture content was determined according to AOAC(1990); the weighed tissue samples were dried at 105 °C in an airoven for 24 h, and then weighed again till a constant weight wasobtained. The protein content was determined using Kjeldahl methodfor determination of nitrogen (Giese, 1967) as follows: % Protein=%nitrogen×6.25 (Giese, 1967; Rigby, 1990). Total lipid content wasdetermined using the Soxhlet extraction method (James, 1995).Carbohydrate was determined according to the method described byJames (1995) using the following equation:

Carbohydrate % = 100−ðmoisture % + protein % + lipid % + ash %Þ:

Ash was determined according to Pearson (1981).

3. Results

3.1. Prey selection and predation rate

The preference of three size groups of H. parthenopeia for fivespecies of bivalve is presented in Table 1. During 30 days, P. undulatawas the most selected prey of the five prey species and it representedabout 70% of the worm's diet, while G. pectinatum were rejectedcompletely. The bivalve C. glaucumwas ranked in the second order asit formed about 20–21.1% of the total prey consumed by the three sizegroups. All three groups of H. parthenopeia appeared to selectapproximately equal numbers of V. pullastra and R. decussata, butthey represented only 3.3–5.5% of the total prey attacked by differentgroups of H. parthenopeia. Results of average daily predation rate ofdifferent size group of H. parthenopeia indicated that large wormsconsumed greater numbers of prey. The highest average dailypredation rates were recorded in case of P. undulata (1.73, 2.13 and2.57 individuals per predator per day for small, medium and largesized groups respectively) followed by C. glaucum (0.50, 0.63 and 0.73individuals per predator per day for the small, medium and large sizedgroups). Meanwhile, the average daily predation rate on V. pullastra

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Table 2Mean values of five bivalves eaten by 3 sized groups of Halla parthenopeia in alternativetrials, presented in pairs during 10 days (values are the means±SD of three replicategroups of 3 worms each).

Prey Predatorsize

Predation rate

Species Shell length (cm) Total number ±SD

Paphia undulata b3.0 Small 174 14.63.0–3.5 Medium 183 12.3N3.5 Large 194 15.5

Cerastoderma glaucum b2.2 Small 12 2.32.2–2.6 Medium 21 3.7N2.6 Large 36 5.5

Paphia undulata b3.0 Small 160 8.13.0–3.5 Medium 171 8.4N3.5 Large 181 15.6

Venerupis pullastra b1.4 Small 8 1.31.4–1.8 Medium 10 2.8N1.8 Large 7 2.3

Paphia undulata b3.0 Small 177 10.33.0–3.5 Medium 187 7.8N3.5 Large 207 13.8

Ruditapes decussata b1.4 Small 7 2.11.4–1.8 Medium 5 1.1N1.8 Large 11 1.9

Paphia undulata b3.0 Small 182 18.93.0–3.5 Medium 188 18.1N3.5 Large 219 22.5

Gafrarium pectinatum b1.8 Small 0 01.8–2.2 Medium 0 0N2.2 Large 0 0

Cerastoderma glaucum b2.2 Small 66 112.2–2.6 Medium 81 5.6N2.6 Large 105 6.6

Venerupis pullastra b1.4 Small 21 8.51.4–1.8 Medium 33 3.4N1.8 Large 30 4.5

Cerastoderma glaucum b2.2 Small 63 6.22.2–2.6 Medium 54 6.9N2.6 Large 57 5.2

Ruditapes decussata b1.4 Small 34 9.61.4–1.8 Medium 39 8.6N1.8 Large 51 6.7

Cerastoderma glaucum b2.2 Small 105 5.32.2–2.6 Medium 134 16.1N2.6 Large 141 7.3

Gafrarium pectinatum b1.8 Small 0 01.8–2.2 Medium 0 0N2.2 Large 0 0

Venerupis pullastra b1.4 Small 33 3.61.4–1.8 Medium 54 5.7N1.8 Large 57 4.8

Ruditapes decussata b1.4 Small 28 4.91.4–1.8 Medium 17 8.3N1.8 Large 27 7.1

Venerupis pullastra b1.4 Small 88 8.71.4–1.8 Medium 84 3.6N1.8 Large 96 2.2

4 I.H. Osman et al. / Journal of Experimental Marine Biology and Ecology 383 (2010) 1–7

and R. decussatawere relatively low (b0.3 individual per predator perday) for all worm size groups (Table 1 and Fig. 2).

Results of feeding preference observations for the three sized groupsofH. parthenopeia, in the trials with the two alternative prey species, arepresented in Table 2. The data in this table confirm the results obtainedin the first experiment. In almost all of the trials one of the prey specieswas consumed in marked preference to the other. Generally, thebivalves P. undulata andC. glaucum appear to be themost preferred foodfor H. parthenopeia. When C. glaucum was offered with V. pullastra orwith R. decussata, all the different sized groups markedly preferredC. glaucum; but when it was offered with P. undulata, all the differentsized groups obviously preferred P. undulata. When V. pullastra andR. decussatawere offered together, V. pullastra seemed more preferableover R. decussata. None of the three sized groups of H. parthenopeiaconsumed any G. pectinatum.

3.2. Growth rate

Table 3 showed the daily growth rate of the three different sizeworm groups when fed with the most two preferred prey species. Forthe two prey species (P. undulata and C. glaucum), growth rate ofH. parthenopeia declines as the worm increases in size. On the otherhand, worms that fed on P. undulata showed higher growth rate thanthat demonstrated by those fed on C. glaucum. During a period of60 days, small sized groups that fed on P. undulata grew from anaverage of 12.9 g to 17.9 g, while medium sized groups grew from18.5 g to 22.8 g and for those large group, worms grew from 23.4 g to25.7 g.

3.3. Feeding behavior

Feeding behavior by H. parthenopeiawas observed throughout theperiod of the experiment. The total period spent during feeding wasdivided into three stages: responding, handling and feeding.Responding to the living food items was considered as the initialstage. During the responding stage, the worm raised its head abovethe sediment and gradually moves in the sediment towards the prey,sometimes the movement was above the sediment and towards thebivalve. Handling starts when the worm first touches the prey to theconsumption of its flesh. Tracks of bivalve movements were observedmany times, which may indicate either escape by the prey or thenormal activity and movement of the bivalve. During feeding stage,the worm wound itself round the bivalve. The bivalve was coveredwith jelly-like material secreted by the polychaete. When the shell ofthe bivalve slightly opens, H. parthenopeia enters the bivalve and eatsall tissue inside the shell. The jelly-like material was largest at the endof feeding along the empty shell.

Fig. 2. Mean daily predation rates±SE of various size groups of Halla parthenopeiaunder laboratory conditions, as the number of prey eaten per predator.

Gafrarium pectinatum b1.8 Small 0 01.8–2.2 Medium 0 0N2.2 Large 0 0

Ruditapes decussata b1.4 Small 71 4.71.4–1.8 Medium 90 6.4N1.8 Large 84 4.4

Gafrarium pectinatum b1.8 Small 0 01.8–2.2 Medium 0 0N2.2 Large 0 0

3.4. Biochemical composition

The percentage composition of moisture content, protein, lipids,carbohydrates and ash of three sized groups of H. parthenopeia areshown in Fig. 3. The moisture content was expressed as percentagewet weight of the tissue, while the biochemical contents weredetermined and expressed as a percentage dry weight of the wholebody. The results showed that there was a high percentage water

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Table 3Daily growth rate of the three size groups of Halla parthenopeia fed with the most two preferred prey species.

Prey Predator

Species Shell length (cm) Soft tissue weight (g) Size Initial weight (g) Final weight (g) Growth rate (g/60 days) Growth rate (g/day)

Paphia undulata b3.0 2.13 Small 12.9 17.9 5.0 0.0833.0–3.5 1.00 Medium 18.5 22.8 4.3 0.072N3.5 0.55 Large 23.4 25.7 2.3 0.038

Cerastoderma glaucum b2.2 0.58 Small 12.6 15.3 2.7 0.0452.2–2.6 0.86 Medium 18.5 20.5 2.0 0.033N2.6 1.26 Large 23.7 24.9 1.2 0.020

5I.H. Osman et al. / Journal of Experimental Marine Biology and Ecology 383 (2010) 1–7

content — more than 70% — for the three size groups and the highestpercentage was recorded in the small size group (average: 76.31%±0.89). Protein was the highest biochemical constituent with anaverage of 51%±2.98 of dry weight, followed by lipids 25.88%±2.76 and carbohydrates 20.72%±3.43. Protein percentage increasedwith increasing weight, and reached the highest in the large sizegroup (average: 53.44%±2.36 dry weight). Lipid content showed aquite similar trend, the percentage was the highest in the large sizedgroup (average: 26.99%±4.30 dry weight). The smallest values forcarbohydrates 17.27%±2.77 and ash contents 2. 3 %±0.12 wererecorded for the large sized groups.

4. Discussion

4.1. Prey selection and growth trials

Carnivorous polychaetes are known to predate on other liveanimals using specialized buccal apparatus. For example, small piecesof fish were eaten by Eurythoe complanata (Pardo and Amaral, 2006),and Onuphid polychaete such as Astralonuphis are known to emergewhere waves break to seek their small sized preys. The feedingbiology of Oenonidae is largely unknown, although they are generallyregarded as carnivores (Rouse and Pleijel, 2001). While Halla okudaiare known to feed on many types of clams (Saito and Imabayashi,1994).

In the present study, the multiple-choice and two alternative trialsof feeding indicated that the H. parthenopeia strongly preferred theP. undulata to the other five bivalve species. Meanwhile, C. glaucumwasthe most preferred bivalve if P. undulatawas not available. Although nopublished data on feeding habits exist for H. parthenopeia in the SuezCanal, it is commonly believed among commercial collectors that itsexistence linked with the presence of C. glaucum. Based on the presentdata, it can be suggested thatH. parthenopeia is selective, searching andchoosing suitable and profitable prey. Predators tend to select their preyin order to minimize the amount of time they spend foraging or tomaximize the amount of energy they attained per prey (Saito et al.,2004). Halla parthenopeia may prefer to feed on P. undulata for its highenergy content. This hypothesis can be supported by the following facts:1- both bivalve species (preys) are available, 2-Cerastoderma glaucum isknown to have no defense mechanism and cannot close the valves

Fig. 3. Average percentages of different biochemical components of three size group ofHalla parthenopeia collected from Suez Canal.

tightly (Nieleson, 1975), which make it easier to be attacked byH. parthenopeia than P. undulata, and 3- Paphia undulata are propor-tionally larger (maximum length N3 cm) than C. glaucum (maximumlength 3 cm) (Gabr, 1991) which may imply higher energy content.However, that H. parthenopeia prefers C. glaucum to V. pullastra andR. decussata may be due to easier handling. Therefore, it can beconcluded that it would be highly beneficial for H. parthenopeia to feedon P. undulata whenever possible. This agrees with Saito et al. (2003),who stated that the shift in prey preference by the polychaetes isprobably limited by constant prey profitability.

Halla parthenopeia did not prey on G. pectinatum either inmultiple-choice trials or in the two alternative trials. This is may beattributed to its defense mechanism, or pheromones from prey as wasdocumented by many authors (Atema, 1995; Chivers and Smith,1998; Nishizaki and Ackerman, 2005). The defense mechanism ofbivalves is governed by the thickness and hardness of the shell, theability of the bivalve to close the valves tightly by the adductormuscles (Thorpe, 1956).

There are very few studies on predator–prey relationship betweenpolychaetes and bivalves (Saito et al., 1999), although polychaetes arecommonly found in sandy intertidal flats. Many other studies havedealt with foraging strategy for other carnivorous organisms such asasteroiods, gastropods and crustaceans (Hughes and Dunkin, 1984;Hughes and Seed, 1995), such predators were suggested to select thediet to maximize their growth (Hughes, 1980).

The growth in marine invertebrates is known to vary withdifferent factors such as temperature, prey and reproductive condi-tion (Ohtomi and Shimizu, 1989; Christou and Verriopoulos, 1993). Inthe present study, the temperature was constant at 25 °C and thespawning season (December–January) (Osman, 2007) differed fromthe period of collection (June), thus the temperature and thereproductive condition seemed to have little effect on growth ofworms and the growth rate of H. parthenopeia was highly affected bythe prey species. This species recorded higher growth rate when feedon more preferred food item (P. undulata) than upon less preferredone (C. glaucum).

The correspondence of prey preference with growth efficiencyindicates that the existence of selective foraging behavior maximizesthe growth (Saito et al., 1999). Considering that the growth of bivalve-feeder was mainly determined by the trade-off between energyintakes versus foraging cost. There are two potential processes of theprey preference to maximize the growth, 1- as with the bivalve-feeding sea star Astropecten articulatus, the predator selects prey withhigher energy content (Beddingfield and McClintock, 1993), and 2- asthe bivalve-feeding gastropod Nucella lapillus, the predator selectsprey with lower handling cost (Burrows and Hughes, 1990).

On the other hand, small sized worms in this study attained highergrowth rate than larger ones. The attainment of higher growth rate isnot for a priority in mature animals, but it may be a vital target forlittle ones. Small animals aremore vulnerable (Janzen et al., 2000) andhave less chance of survival when foraging (Hughes and Seed, 1995).Another species of the genusHalla, H. okudai, has been studied well bySaito et al. (1999) to show the relationship between its growthand species and abundance of prey, they found that the growth of

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H. okudai increased by feeding on the more preferred prey with lowsearching cost (R. philipppinarum) rather than the less preferred(Crassostrea gigas) with higher searching cost.

4.2. Feeding behavior

Results of the present study revealed that, the most suitablesediment composition for living of H. parthenopeia had about 50%mud fraction, which differed from the soil mixture of H. okudai(Imabayashi et al., 1996). The highly gravel-low mud sediment for H.okudaiwas not favored by H. parthenopeia in the present study and inthe preliminary trials; worms frequently could not burrow into thesediment and died few days after the beginning of the experiment.

The behavior by H. parthenopeia when foraging has not beendocumented before, but observations in this study were similar tothat of H. okudai (Imabayashi et al., 1996). It was observed thatH. parthenopeia secretes a large amount of jelly-like substancesthrough handling and feeding upon prey bivalves. The secretion ofthis slimy material was not restricted to feeding only, but was alsosecreted during handling and touching of the worm (personalobservation), and when the worm was wounded, although thissecretion was highly aggregated, it differed in transparency from thatproduced during feeding. However, no information has been compiledyet concerning the biochemical characters of the jelly-like materialsecreted while handling or feeding on prey.

Secretion of jelly-like material during feeding is a peculiar feedingstyle to polychaetes and it was documented by many polychaetesfrom different families. Halla okudai, an oenonid polychaete, has welldeveloped pharyngeal jaws and feed on clams by aiding of jelly-likematerial during handling and feeding of prey (Kawai et al., 1999).Arabella iricolor is a polychaete species from Arabellidae; it has beenknown to feed on the mussel Mytilis edulis while secreting a slimysubstance (Iwasaki, 1997).

4.3. Biochemical composition

Very few published papers are available for comparisons ofbiochemical composition of Oenonid polychaetes. In this study thebiochemical composition of H. parthenopeia showed that protein wasthe highest biochemical constituent accounting for an average of 51%of the dry weight, followed by lipids with an average of 25.88%,meanwhile carbohydrate was present at relatively good level with anaverage of 20.72%. Lipids are very important in the physiology ofmarine animals, as has been stressed by Giese (1966) and Morris andSargent (1973). Marine animals rely to a larger extent than mostterrestrial ones on lipids rather than on carbohydrate for metabolicenergy reserves (Cowey and Sargent, 1972; Holland, 1978). Inaddition, the protein is the major dietary component that influencesthe growth of different cultured organisms (Robinson and Li, 1999;Singh et al., 2006). The present results indicated that the highpercentage content of protein and lipids of H. parthenopeia lend it tobeing a good candidate for the successful production of high qualityjuveniles in both finfish and crustaceans.

In conclusion, this study provides the first known account onfeeding behavior, growth and biochemical composition of the mostcommercially feasible polychaete species in Suez Canal. The results ofthis study indicated that H. parthenopeia are capable of using manydifferent types of clams, although they grew best on P. undulata in thepresence of 50% clay sediment. Halla parthenopeia is rich in proteinand lipid and would be of highly nutritious if used as feed in fishculture especially for feeding brood stock. Although these results arepreliminary, our findings suggest the feasibility of successful cultureof this species in captivity. The elucidation of these aspects and otherquestions regarding to culture of this species will make it possible, inthe near future, to develop large-scale artificial systems. Large-scaleculture provides not only an increasing market of live bait, but will

also reduce the substrate harvesting disturbance, and decrease thebiological impact on benthic communities, with lower environmentaleffects (Olive, 1994).

Acknowledgements

We would like to thank Dr. Maher A. Aziz, the lecturer in MarineScience Department, for his great help in preparation of the aquaria.His kind assistance and relief during maintaining the worms in theirnew home are greatly appreciated. This study is part of the M.Scdissertation in Marine Biology, Suez Canal University. [SS]

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